Insectivorous Plants

(1) A fourth bladder contained an articulate organism, for I distinctly saw the remnant of a limb, terminating in a hook. The quadrifids were not examined. (5) A fifth included much decayed matter apparently of some animal, but with no recognisable features. The quadrifids in contact contained numerous spheres of protoplasm. (G) Some few bladders on the plant which I received from Kew were examined ; and in one, there was a worm-shaped animal very little decayed, with a distinct remnant of a similar one greatly decayed. Several of the arms of the processes in contact with these remains contained two spherical masses, like the single solid nucleus which is properly found in each arm. In another bladder there was a minute grain of quartz, remind- ing me of two similar cases with Utric.ularia neglecta.

As it appeared probable that this plant would capture a greater number of animals in its native country than under culture, I obtained permission to remove small portions of the rhizomes from dried specimens in the herbarium at Kew. I did not at first find out that it was advisable to soak the rhizomes for two or three days, and that it was necessary to open the bladders and spread out their contents on glass ; as from their state of decay and from having been dried and pressed, their nature could not otherwise be well distinguished. Several bladders on a plant which had grown in black earth in New Granada were first examined ; and four of these included remnants of animals. The first contained a hairy Acarus, so much decayed that notliing was left except its transparent coat

also a yellow chitinous head of some animal with art internal fork, to which the oasophagus was suspended, but I could see no mandibles; also the double hook of the tarsus of some animal ; also an elongated greatly decayed animal ; and lastly, a curious flask-shaped organism, having the walls formed of rounded cells. Professor Glaus has looked at this latter organism, and thinks that it is the shell of a rhizopod, probably one of the Arcellidfe. In this bladder, as well as in several others, there were some unicellular Algae, and one multicellular Alga, which no doubt had lived as intruders.

A second bladder contained an Acarus much less decayed than the former one, with its eight legs preserved ; as well as remnants of several other articulate animals. A third bladder contained the end of the abdomen with the two hinder limbs of an Acarus, as I believe. A fourth contained remnants of a distinctly articulated bristly animal, and of several other organ- isms, as well as much dark brown organic matter, the nature of which could not be made out.

Some bladders from a plant, which had lived as an epiphyte iu Trinidad, in the West Indies, were next examined, but not so carefully as the others; nor had they been soaked long enough. Four of them contained much brown, translucent, granular matter, apparently organic, but with no distinguish- able parts. The quadrifids iu two were brownish, with their contents granular ; and it was evident that they had absorbed matter. In a fifth bladder there was a flask-shaped organism, like that above mentioned. A sixth contained a very long, much decayed, worm-shaped animal. Lastly, a seventh bladder contained an organism, but of what nature could not be dis- tinguished.

Only one experiment was tried on the quadrifid pro- cesses and glands with reference to their power of absorption. A bladder was punctured and left for 24 hrs. in a solution of one part of urea to 437 of water, and the quadrifid and bifid processes were found much affected. In some arms there was only a single symmetrical globular mass, larger than the proper nucleus, and consisting of yellowish matter, generally translucent but sometimes granular; in others there were two masses of different sizes, one large and the

other small; and in others there were irregularly shaped globules ; so that it appeared as if the limpid contents of the processes, owing tc the absorption of matter from the solution, had become aggregated sometimes round the nucleus, and sometimes into sepa- rate masses; and that these then tended to coalesce. The primordial utricle or protoplasm lining the pro- cesses was also thickened here and there into irregular and variously shaped specks of yellowish translucent matter, as occurred in the case of Utrieularia neglecta under similar treatment. These specks apparently did not change their forms.

The minute two-armed glands on the valve were also affected by the solution ; for they now contained several, sometimes as many as six or eight, almost spherical masses of translucent matter, tinged with yellow, which slowly changed their forms and posi- tions. Such masses were never observed in these glands in their ordinary state. We may therefore infer that they serve for absorption. Whenever a little water is expelled from a bladder containing animal remains (by the means formerly specified, more especially by the generation of bubbles of air), it will fill the cavity in which the valve lies ; and thus the glands will be able to utilise decayed matter which otherwise would have been wasted.

Finally, as numerous minute animals are captured by this plant in its native country and when culti- vated, there can be no doubt that the bladders, though so small, are far from being in a rudimentary con- dition ; on the contrary, they are highly efficient traps. Nor can there be any doubt that matter is absorbed from the decayed prey by the quadrifid and bifid processes, and that protoplasm is thus generated. What tempts animals of such diverse kinds to enter

the cavity beneath the bowed antennae, and then force their way through the little slit-like orifice between the valve and collar into the bladders filled with water, I cannot conjecture. Tubers. — These organs, one of which is represented in a previous figure (fig. 26) of the natural size, deserve a few remarks. Twenty were found on the rhizomes of a single plant, but they cannot be strictly counted ; for, besides the twenty, there were all pos- sible gradations between a short length of a rhizome just perceptibly swollen and one so much swollen that it might be doubtfully called a tuber. When well developed, they are oval and symmetrical, more so than appears in the figure. The largest which I saw was 1 inch (25'4 mm.) in length and '45 inch (11-43 mm.) in breadth. They commonly lie near the surface, but some are buried at the depth of 2 inches. The buried ones are dirty white, but those partly exposed to the light become greenish from the development, of chlorophyll in their superficial cells. They terminate in a rhizome, but this sometimes decays and drops off. They do not contain any air, and they sink in water; their surfaces are covered with the usual papillse. The bundle of vessels which runs up each rhizome, as soon as it enters the tuber, separates into three distinct bundles, which reunite at the opposite end. A rather thick slice of a tuber is almost as transparent as glass, and is seen to consist of large angular cells, full of water and not containing starch or any other solid matter. Some slices were left in alcohol for several days, but only a few extremely minute granules of matter were precipitated on the walls of the cells ; and these were much smaller and fewer than those precipitated on the cell-walls of the rhizomes and bladders. We may therefore con-

elude that the tubers do not serve as reservoirs lor food, but for water during the dry season to which the plant is probably exposed. The many little bladders filled with water would aid towards the same end. To test the correctness of this view, a small plant, growing in light peaty earth in a pot (only 4-i- by 4-^ inches outside measure) was copiously watered, and then kept without a drop of water in the hothouse. Two of the upper tubers were beforehand uncovered and measured, and then loosely covered up again. In a fortnight's time the earth in the pot appeared ex- tremely dry ; but not until the thirty-fifth day were the leaves in the least affected ; they then became slightly reflexed, though still soft and green. This plant, which bore only ten tubers, would no doubt have resisted the drought for even a longer time, had I not previously removed three of the tubers and cut off several long rhizomes. When, on the thirty-fifth day, the earth in the pot was turned out, it appeared as dry as the dust on a road. All the tubers had their surfaces much wrinkled, instead of being smooth and tense. They had all shrunk, but I cannot say accurately how much ; for as they were at first symmetrically oval, I measured only their length and thickness ; but they contracted in a transverse line much more in one direction than in another, so as to become greatly flattened. One of the two tubers which had been measured was now three-fourths of its original length, and two- thirds of its original thick- ness in the direction in which it had been measured, but in another direction only one-third of its former thickness. The other tuber was one-fourth shorter, one- eighth less thick in the direction in which it had been measured, and only half as thick in another direction.

and examined. The cells still contained much water and no air, but they were more rounded or less angular than before, and their walls not nearly so straight ; it was therefore clear that the cells had contracted. The tubers, as long as they remain alive, have a strong attraction for water ; the shrivelled one, from which a slice had been cut, was left in water for 22 hrs. 30 m., and its surface became as smooth and tense as it originally was. On the other hand, a shrivelled tuber, which by some accident had been separated from its rhizome, and which appeared dead, did not swell in the least, though left for several days in water.

With many kinds of plants, tubers, bulbs, &c. no doubt serve in part as reservoirs for water, but I know of no case, besides the present one, of such organs having been developed solely for this purpose. Prof. Oliver informs me that two or three other species of Utricularia are provided with these appendages ; and the group containing them has in consequence received the name of orchidioides. All the other species of Utricularia, as well as of certain closely related genera, are either aquatic or marsh plants; therefore, on the principle of nearly allied plants generally having a similar constitution, a never failing supply of water would probably be of great importance to our present species. We can thus understand the meaning of the development of its tubers, and of their number on the same plant, amounting in one instance to at least twenty.

As I wished to ascertain whether the bladders on the rhizomes of other species of Utricularia, and of the species of certain closely allied genera, had the same essential structure as those of Utricularia montana, and whether they captured prey, I asked Prof. Oliver to send me fragments from the herbarium at Kew. He kindly selected some of the most distinct forms, having entire leaves, and believed to inhabit marshy ground or water. My son, Francis Darwin, examined them, and has given me the following observations; but it should be borne in mind that it is extremely difficult to make out the structure of such minute and delicate objects after they have been dried and pressed.*

Utricularia nelunibifulia (Organ Mountains, Brazil). — The habitat of this species is remarkable. According to its discoverer, Mr. Gardner,! it is aquatic, but " is only to be found growing in the water which collects in the bottom of the leaves of a large Tillandsia, that inhabits abundantly an arid rocky part of the moun- tain, at an elevation of about 5000 feet above the level of the sea. Besides the ordinary method by seed, it propagates itself by runners, which it throws out from the base of the flower-stem ; this runner is always found directing itself towards the nearest Tillandsia, when it inserts its point into the water and gives origin to a new plant, which in its turn sends out another shoot. In this manner I have seen not less than six plants united." The bladders resemble those of Utricularia montana in all essential respects, even to the presence of a few minute two-armed glands on the valve. Within one bladder there was the remnant of the abdomen of some larva or crustacean of large size,

* Prof. Oliver has given (' Proc. but he does not appear to have Linn. Soc.' vol. iv. p. 169) figures paid particular attention to these American species, namely, Utri- f 'Travels in the Interior of having a brush of long sharp bristles at the apex. Other bladders included fragments of articulate ani- mals, and many of them contained broken pieces of a curious organism, the nature of which was not recog- nised by anyone to whom it was shown. Utricularia amethystina (Guiana). — This species has small entire leaves, and is apparently a marsh plant; but it must grow in places where crustaceans exist, for there were two small species within one of .the bladders. The bladders are nearly of the same shape as those of Utricularia montana, and are covered outside •with the usual papillae ; but they differ remarkably in the antennas being reduced to two short points, united by a membrane hollowed out in the middle. This membrane is covered with innumerable oblong glands supported on long footstalks ; most of which are arranged in two rows converging towards the valve. Some, however, are seated on the margins of the mem- brane ; and the short ventral surface of the bladder, between the petiole and valve, is thickly covered with glands. Most of the heads had fallen off, and the foot- stalks alone remained ; so that the ventral surface and the orifice, when viewed under a weak power, appeared as if clothed with fine bristles. The valve is narrow, and bears a few almost sessile glands. The collar against which the edge shuts is yellowish, and presents the usual structure. From the large number of glands on the ventral surface an-,1 round the orifice, it is probable that this species lives in very foul water, from which it absorbs matter, as well as from its captured and decay- ing prey.

Utricularia griffitliii (Malay and Borneo). — The bladders are transparent and minute ; one which was measured being only t%»0 of an inch ('711 mm.) in diameter. The antennae are of moderate length, and project straight forward ; they are united for a short space at their bases by a membrane ; and they bear a moderate number of bristles or hairs, not simple as heretofore, but surmounted by glands. The bladders also differ remarkably from those of the previous speciesj as within there are no quadrifid, only bifid, processes. In one bladder there was a minute aquatic larva; in another the remains of some articulate animal ; and in most of them grains of sand.

Utricularia cserulea (India). — The bladders re- semble those of the last species, both in the general character of the antennae and in the processes with- in being exclusively bifid. They contained remnants of entomostracan crustaceans, Utricularia orbiculata (India). — The orbicular leaves and the stems bearing the bladders apparently float in water. The bladders do not differ much from those oJ the two last species. The antennae, which are united for a short distance at their bases, bear on their outer surfaces and summits numerous, long, multicellular hairs, surmounted by glands. The processes within the bladders are quadrifid, with the four diverging arms of equal length. The prey which they had captured consisted of entomostracan crustaceans.

Utricularia multicaulit, (Sikkim, India, 7000 to 11,000 feet). - -The bladders, attached to rhizomes, are remarkable from the structure of the antennae. These are broad, flattened, and of large size; they bear on their margins multicellular hairs, surmounted by glands. Their bases are united into a single, rather narrow pedicel, and they thus appear like a great digitate expansion at one end of the bladder. Internally the quadrifid processes have divergent arms of equal length. The bladders contained remnants of articulate animals.

This genus, which is confined to Western Australia, is characterised by having a " quadripartite calyx." In other respects, as Prof. Oliver remarks,* " it is quite a Utricularia." Polypompliolyx muUifida. — The bladders are attached in whorls round the summits of stiff stalks. The two antennae are represented by a minute membranous fork, the basal part of which forms a sort of hood over the orifice. This hood expands into two wings on each side of the bladder. A third wing or crest appears to be formed by the extension of the dorsal surface of the petiole ; but the structure of these three wings could not be clearly made out, owing to the state of the speci- mens. The inner surface of the hood is lined with long simple hairs, containing aggregated matter, like that within the quadrifid processes of the previously described species when in contact with decayed ani- mals. These hairs appear therefore to serve as absor- bents. A valve was seen, but its structure could not be determined. On the collar round the valve there are in the place of glands numerous one-celled papillae, having very short footstalks. The quadrifid processes have divergent arms of equal length. Eemains of entomostracan crustaceans were found within the bladders.

Polypomplwlyx tenella. — The bladders are smaller than those of the last species, but have the same general structure. They were full of debris, apparently organic, but no remains of articulate animals could be distinguished. This remarkable genus is technically distinguished from Utricularia, as I hear from Prof. Oliver, by having a five-partite calyx. Species are found in several parts of the world, and are said to be " herbae annure paludosoe."

Genlisea ornata (Brazil). — This species has been described and figured by Dr. Warming,* who states that it bears two kinds of leaves, called by him spathulate and utriculiferous. The latter include cavities ; and as these differ much from the bladders of the foregoing species, it will be convenient to speak of them as utricles. The accompanying figure (fig. 29) of one of the utriculiferous leaves, about thrice en- larged, will illustrate the following description by my son, which agrees in all essential points with that given by Dr. Warming. The utricle (b) is formed by a slight enlargement of the narrow blade of the leaf. A hollow neck (n), no less than fifteen times as long as the utricle itself, forms a passage from the transverse slit-like orifice (o) into the cavity of the utricle. A utricle which measured TV of an inch (•705 mm.) in its longer diameter had a neck -i-f (10-583 mm.) in length, and -^ of an inch ('254 mm.) in breadth. On each side of the orifice there is a long spiral arm or tube (a) ; the structure of which will be best understood by the following illustration. Take a narrow ribbon and wind it spirally round a thin cylinder, so that the edges come into contact along its whole length; then pinch up the two edges so as to form a little crest, which will of course wind spirally

"Bidrag til Kundskaben om Lentibulariacese," Copenhagen, 1874 lower part of the utricle is covered with spherical papillae, formed of four cells (sometimes eight accord- ing to Dr. Warming), which evidently answer to the quadrifid processes within the bladders of Utricularia. I Upper part of lamina of leaf. 6 Utricle or bladder. n Neck of utricle. o Orifice. These papillae extend a little way up the dorsal and ventral surfaces of the utricle ; and a few, according to Warming, may be found in the upper part. This upper region is covered by many transverse rows, one above the other, of short, closely approximate hairs, pointing downwards. These hairs have broad bases,

and their tips are formed by a separate cell. They are absent in the lower part of the utricle where the pa- pillre abound. The neck is likewise lined throughout its whole length with trans- verse rows of long, thin, transparent hairs, having broad bulbous (fig. 30) bases, with similarly constructed sharp points. They arise from little projecting ridges, formed of rectangular epi- dermic cells. The hairs vary a little in length, but their points generally extend down to the row next below ; so that if the neck is split open and laid flat, the inner surface re-

portion of inside of neck leading the hairs representing the ing the downward pointed bristles, pins, and the little transverse pins are thrust. These rows of hairs are indicated in the previous figure (29) by numerous transverse lines crossing the neck. The inside of the neck in also studded with papillae ; those in the lower part are spherical and formed of four cells, as in the lower part of the utricle ; those in the upper part are formed of two cells, which are much elongated downwards beneath their points of attachment. These two-celled papillse apparently correspond with the bifid process in the upper part of the bladders of Utricularia. The narrow transverse orifice (o, fig. 29) is situated between the bases of the two spiral arms. No valve could be detected here, nor was any such structure seen by Dr. Warming. The lips of the orifice are armed with many short, thick, sharply pointed, somewhat incurved hairs or teeth.

The two projecting edges of the spirally wound lamina, forming the arms, are provided with short incurved hairs or teeth, exactly like those on the lips. These project inwards at right angles to the spiral line of junction between the two edges. The inner surface of the lamina supports two-celled, elon- gated papillae, resembling those in the upper part of the neck, but differing slightly from them, according to Warming, in their footstalks being formed by prolongations of large epidermic cells ; whereas the papillae within the neck rest on small cells sunk amidst the larger ones. These spiral arms form a conspicuous difference between the present genus and Utricularia.

Lastly, there is a bundle of spiral vessels which, "running up the lower part of the linear leaf, divide? close beneath the utricle. One branch extends up the dorsal and the other up the ventral side of both the utricle and neck. Of these two branches, one enters one spiral arm, and the other branch the other arm. The utricles contained much debris or dirty matter, which seemed organic, though no distinct organisms could be recognised. It is, indeed, scarcely possible that any object could enter the small orifice and pass down the long narrow neck, except a living creature. Within the necks, however, of some specimens, a worm with retracted horny jaws, the abdomen of some articulate animal, and specks of dirt, probably the remnants of other minute creatures, were found. Many of the papillae within both the utricles and necks were discoloured, as if they had absorbed matter. From this description it is sufficiently obvious how Genlisea secures its prey. Small animals entering the narrow orifice — but what induces them to enter is not known any more than in the case of Utricularia — would find their egress rendered difficult by the sharp incurved hairs on the lips, and as soon as they passed some way down the neck, it would be scarcely possible for them to return, owing to the many transverse rows of long, straight, downward pointing hairs, together with the ridges from which these project. Such crea- tures would, therefore, perish either within the neck or utricle ; and the quadrifid and bifid papillae would absorb matter from their decayed remains. The transverse rows of hairs are so numerous that they seem superfluous merely for the sake of preventing the escape of prey, and as they are thin and delicate, they probably serve as additional absorbents, in the same manner as the flexible bristles on the infolded margins of the leaves of Aldrovanda. The spiral arms no doubt act as accessory traps. Until fresh leaves are examined, it cannot be told whether the line of junction of the spirally wound lamina is a little open along its whole course, or only in parts, but a small creature which forced its way into the tube at any point, would be prevented from escaping by the incurved hairs, and would find an open path down

the tube into the neck, and so into the utricle. If the creature perished within the spiral arms, its decaying remains would be absorbed and utilised by the bifid papillae. We thus see that animals are captured by Genlisea, not by means of an elastic valve, as with the foregoing species, but by a contrivance resembling an eel-trap, though more complex. Genlisea africana (South Africa). — Fragments of the utriculiferous leaves of this species exhibited the same structure as those of Genlisea ornata. A nearly perfect Acarus was found within the utricle or neck of one leaf, but in which of the two was not recorded.

Genlisea aurea (Brazil). — A fragment of the neck of a utricle was lined with transverse rows of hairs, and was furnished with elongated papillae, exactly like those within the neck of Genlisea ornata. It is probable, therefore, that the whole utricle is similarly constructed. Genlisea fili/ormis (Bahia, Brazil). — Many leaves were examined and none were found provided with utricles, whereas such leaves were found without diffi- culty in the three previous species. On the other hand, the rhizomes bear bladders resembling in essen- tial character those on the rhizomes of Utricularia. These bladders are transparent, and very small, viz. only -pi-o of an inch ('254 mm.) in length. The antennae are not united at their bases, and apparently bear some long hairs. On the outside of the bladders there are only a few papillae, and internally very few quadrifid processes. These latter, however, are of un- usually large size, relatively to the bladder, with the four divergent arms of equal length. No prey could be seen within these minute bladders. As the rhizomes of this species were furnished with bladders, those of Genlisea africana, ornata, and aurea were carefully

examined, but none could be found. What are we to infer from these facts? Did the three species just named, like their close allies, the several species of Utricularia, aboriginally possess bladders on their rhizomes, which they afterwards lost, acquiring in their place utriculiferous leaves ? In support of this view it may be urged that the bladders of Genlisea filiformis appear from their small size and from the fewness of their quadrifid processes to be tending towards abortion ; but why has not this species acquired utriculiferous leaves, like its congeners ?